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Ernodesmis and Apjohnia Phycologia (1975) Volume 14 (4), 309-316 The morphology and systematic position of the green algae Ernodesmis and Apjohnia GEORGE F. PAPENFUSS AND Mnsuo CHIHARA Department of Botany, University of California, Berkeley, California 94720, U.S.A. and Department of Botany, Tokyo Kyoiku University, 3-29-1, Otsuka, Bunkyo-ku, Tokyo 112, Japan The monotypic genera Ernodesmis B0rgesen and Apjohnia Harvey, which in habit resemble each other and which have been regarded as belonging to the Siphonocladaceae by most authors, are shown to be different genera. Neither appears to belong to the Siphonocladaceae. Ernodesmis fits well in the Valoniaceae (Siphonocladales) and Apjohnia is provisionally assigned to the Anadyomenaceae (Cladophorales). Peculiar intracellular rhizoid-like cells are produced at the base of branches in older parts of the thallus of Apjohnia. Comparable structures are also formed in several other green algae, including Ernodesmis and Cladophora rugulosa. They are believed to give mechanical strength at the juncture of adjacent cells, especially at the place of insertion of branches. Introduction Morphology From published descriptions and illustrations of B0rgesen (1912, 1913) has fully described and Ernodesmis and Apjohnia, the impression is gained illustrated the development of the thallus; a brief that these two genera of marine green algae may be account will suffice, therefore. The plant starts as congeneric. The present study was undertaken with an erect, clavate, vesicular cell, 10-20 mm long and the hope of resolving this problem. 1·5-2·5 mm in diameter, with basal annulations, attached by a system of septate rhizoids. This cell produces a terminal cluster of five to eight or more Ernodesmis Borgesen one-celled branches, each separated by a concave wall from the parent cell. Each of these primary History of the genus branches produces in turn a similar terminal whorl Ernodesmis was erected by B0rgesen in 1912 (see of branches (Figs 1-3). This process is repeated also B0rgesen, 1913, pp. 66-71) for a species from until six (or more?) orders of branches have been St Croix, Virgin Islands that Kiitzing had described formed. The mature plant attains a height of about in 1849 and illustrated in 1856 (pI. 88) under the 5 cm. Not infrequently, hapteroid rruzoidal cells are name Valonia verticil/ata. The plant is widely produced at the proximal end of the cells (see also distributed in the Caribbean (Taylor, 1960) and is B0rgesen, 1912, 1913). Similar rhizoids were called also known from Brazil (Schmidt, 1924), the Canary clamps by Murray & Boodle (1888, p. 276) with Islands (B0rgesen, 1925), several places in West reference to Struvea ramosa. They elongate and Africa (Steen toft, 1967), the west coast of Mexico attach themselves firmly to the distal end of the cell (Dawson, 1961), including the northern part of the below. Gulf of California (Dawson, 1966), Ceylon (Dura i­ The cells are multinucleate and contain numerous, ratnam, 1961), Mauritius (B0rgesen, 1940, 1946) small, polygonal chloroplasts (each provided with a and Kenya (Gerloff,1960). pyrenoid) joined at the corners to adjacent plastids B0rgesen (1913, 1925) placed Ernodesmis in the to form a reticulum. Siphonocladaceae, in which family it has been The earliest events in the initiation of branches in retained by all authors dealing with it, except Ernodesmis were not observed by us. From slightly Oltmanns (1922), Fritsch (1935) and Taylor (1960), later stages it is seen, however, that a branch always who placed it in the Valoniaceae. Fritsch and Taylor, has its origin as a lenticular cell formed in the however, did not accept the family Siphonoclada­ terminal region of the parent cell (Figs 1 and 3). ceae. This cell produces a protuberance (Fig. 2) which 309 310 Phyco[ogia, Vol. 14 (4), 1975 2 mm \ 2 FIGS 1-3. Ernodesmis verticil/ato. Stages in the development of lateral branches; note the origin of a branch from a lenticular cell (Fig. I) with a concave basal septum. Papenfuss and Chihara: Ernodesmis and Apjohnia 311 elongates and forms a branch with a concave basal the many species of the genus possess annular septum which has been present since the inception constrictions. of the branch (Figs 1-3). Apjo/mia Harvey Discussion History of the genus The method of branch formation in Ernodesmis Apjolmia, with the species A. laetevirens, was is like that described by Papenfuss & Egerod (1957) described by Harvey in 1855 upon material from for Valoniopsis pachynema, and also recalls the Australia (Phillip Island, Western Port, Victoria) situation in Valonia (Egerod, 1952). It is unlike the and in 1858 he illustrated it in the first volume of his situation in Siphonoc/adus in which the branches are Phycologia australica (pI. 5). J. Agardh (1887, pp. endogenous in origin as a result of segregative 105-108) reported the species from Tasmania. division of the protoplast of the parent cell (see According to Womersley (1956), A. laetevirens in Egerod, 1952, fig.2m-o). mainland Australia ranges from Elliston, South Ern.odesmis agrees with Valonia in forming Australia, to Collaroy (near Sydney), New South spherical cytoplasmic masses in some of the cells Wales, whence it was obtained in the drift by May (Fig. 3). These bodies, which were also observed (1946). KUtzing in 1863 (p. 12) described and later by KUtzing (1856) in material of this genus from the (1866, pI. 2) illustrated a plant from New Caledonia Virgin Islands and by B0rgesen (1913, 1940) in (lie des Pins) under the name Struvea scoparia. specimens from the Virgin Islands and Mauritius, Murray & Boodle (1888, pp. 266 and 281) merged are frequently present in Valonia, for which they this species in Apjohnia laetevirens, remarking, were first reported by Murray (1893). They have however, that it was not in perfect accord with the been regarded as asexual reproductive structures, latter. Womersley (1956) is inclined to doubt the produced perhaps under unfavourable environ­ conspecificity of these taxa. Denizot (1965) has mental conditions. Their exact function remained rediscovered KUtzing's plant in New Caledonia and uncertain, however, until it was shown by Chihara has discussed and illustrated it under the name (1959) that they can function as aplanospores and A. laetevirens. In habit, the plant from New Cale­ are capable of developing into new plants. donia (Denizot, 1965, pI. [1]) does not appear to Fritsch (1935, p. 424) was of the opinion that correspond exactly to the one from Australia, but 'Ernodesmis, Siphonoc/adus, Struvea, and Chamaedoris whether this difference is sufficient to reiain it as form another apparently natural group characterized an autonomous species or to justify its removal to by the annular constrictions on the main axes and another genus cannot be settled without examination sometimes also on the branches'. However, the of the plant. formation of annular constrictions at the bases of The brothers Crouan in 1865 (in Schramm & axes (and at times of the branches) apparently is a Maze, p. 47) described a plant from Guadeloupe feature that has arisen several times in the green Island in the West Indies as Apjohnia tropica. J. alga�, and in most instances it probably is not Agardh in 1887 removed this species to Siphono­ indicative of phylogenetic affinity. Thus, for example, c/adlls Schmitz, where it has remained to the present. Struvea anastomosans, S. tenuis and S. pulcherrima Murray (1891, p. 209) transferred the South (Boodleaceae, Siphonocladales) lack annular con­ African Cladophora 1"1Iguiosa Martens (1866) to strictions in the stipe whereas the other species of Apjohnia. Although there are features of agreement the genus possess them (Egerod, 1952, p. 361); between A. laetevirens and C. ruglllosa, the evidence some species of Caulerpa (Caulerpales), e.g. the favours keeping C. rugulosa in the genus Cladophora. South African C. /ili/ormis and C. holmesiana, Wille (1910, p. 111) reduced the monotypic possess annular constrictions in the stipe whereas Rudicularia Heydrich (J 903), based on material the great majority of species of the genus lack them; from the Ryukyu Islands, to a synonym of Apjohnia in the Cladophorales, Chaetomorpha antennina and made the combination A. penicillata (Heydrich). possesses annular constrictions but all the other RlIdiclllaria probably has little, if anything, in com­ species of the genus, to our knowledge, lack them; mon with Apjohnia. It may be congeneric with in Cladophora itself, as far as we are aware, only Tydemania Weber-van Bosse but for the present its C. rugulosa, C. wrightiana and C. prolt/era among identity must be regarded as uncertain. (See Papen- 312 Phy'!;Vlogia, Vol.14 (4),1975 fuss, 1953, p. 30, for a brief discussion of Rudicu­ dendroid tuft to 15 cm high (Harvey, 1858). It is laria.) attached by a system of branched, sparsely septate Harvey (1855), J. Agardh (1887), Murray (1895), rhizoids. The plant starts as an erect, club-shaped Wille (1890, 1910) and Printz (1927) placed Apjohnia vesicular cell, which produces rhizoids at its proximal in the Valoniaceae whereas Oltmanns (1904), end and at its summit a dome-shaped outgrowth Feldmann (1938) and Womersley (1956) placed it in which will become an axial cell. After the dome has the Siphonocladaceae. B0rgesen (1913) and West been formed, a whorl of conical outgrowths with (1916) treated the Siphonocladaceae as a subfamily obtuse apices is produced round the central axial of the Valoniaceae and referred Apjohnia to it. cell (Figs 4 and 5). All of these outgrowths elongate Several authors (Wille, 1890; B0rgesen, 1912 ; considerably and are ultimately separated by basal Printz, 1927) considered Apjohnia to be a nonseptate septa from the lower primary cell, which functions alga, an erroneous belief that caused Setchell (1929) as a stalk cell. The cells surrounding the axial cell to suspect that the genus should be placed in develop into a whorl ofJatera\ branches.
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